Effect of Crop Rotation on Meloidogyne spp. and Pratylenchus spp. Populations in Strawberry Fields in Taiwan

Changes in population levels of Meloidogyne hapla, M. incognita, Pratylenchus coffeae, and P. penetrans were studied in 12 strawberry fields in the Dahu region of Taiwan. Ten potential rotation crops and two cultural practices were evaluated for their effect on nematode populations and influence on strawberry yield. Rotation with rice or taro and the cultural practice of flooding and bare fallowing for four months were found to reduce nematode soil populations to two or fewer nematodes per 100 ml soil. Average strawberry yields increased between 2.4% to 6.3% following taro compared to the bare fallow treatment. Corn suppressed M. incognita and M. hapla populations and resulted in an increased in strawberry yield compared to bare fallow. Other phytopathogens also present in these fields limited taro as the rotation choice for nematode management. Results of this research and economic analysis of the input requirements for various rotation crops, corn and bare fallow were recommended as the most appropriate rotation strategies for nematode management in strawberry in this region.     

Use of Green Manure Crops in Control of Hirschmanniella mucronata and H. oryzae in Irrigated Rice

Four field experiments were conducted to study the effect of Sesbania rostrata and Aeschynomene afraspera as rotational and green manure crops on the population dynamics of Hirschmanniella mucronata and H. oryzae, and subsequent rice yields. The sequential cropping of the legumes with rice controlled both nematode species. In two experiments, yield of rice was related to the nematode population denisites at planting and harvesting of the second rice crop (R² = 0.391, P < 0.001, and R² = 0.57, P < 0.001), regardless of the treatments. Rice yield increases were attributed to nutritional effect of the green manure and the reduction of the nematode populations or the modification of a factor(s) linked to the nematode populations induced by their cropping. As the two leguminous crops do not generate direct return, using them to control the rice-root nematodes was not economical, despite the significant yield increase obtained.     

Sequential Decision Rules for Managing Nematodes with Crop Rotations

A dynamic model of nematode populations under a crop rotation that includes both host and nonhost crops is developed and used to conceptualize the problem of economic control. The steady state of the dynamic system is used to devise an approximately optimal decision policy, which is then applied to cyst nematode (Heterodera schachtii) control in a rotation of sugarbeet with nonhost crops. Long-run economic returns from this approximately optimal decision rule are compared with results from solution of the exact dynamic optimization model. The simple decision rule based on the steady state provides long-run average returns that are similar to the fully optimal solution. For sugarbeet and H. schachtii, the simplified rule can be calculated by maximizing a relatively simple algebraic expression with respect to the number of years in the sequence of nonhost crops. Maximization is easy because only integers are of interest and the number of years in nonhost crops is typically small. Solution of this problem indirectly yields an approximation to the optimal dynamic economic threshold density of nematodes in the soil. The decision rule requires knowledge of annual nematode population change under host and nonhost crops, and the relationship between crop yield and nematode population density.     

Effect of Rotation Crops on Heterodera glycines Population Density in a Greenhouse Screening Study

Crop rotation is a common means of reducing pathogen populations in soil. Several rotation crops have been shown to reduce soybean cyst nematode (Heterodera glycines) populations, but a comprehensive study of the optimal crops is needed. A greenhouse study was conducted to determine the effect of growth and decomposition of 46 crops on population density of H. glycines. Crops were sown in soil infested with H. glycines. Plants were maintained until 75 days after planting, when the soil was mixed, a sample of the soil removed to determine egg density, and shoots and roots chopped and mixed into the soil. After 56 days, soil samples were again taken for egg counts, and a susceptible soybean (‘Sturdy’) was planted in the soil as a bioassay to determine egg viability. Sunn hemp (Crotalaria juncea), forage pea (Pisum sativum), lab-lab bean (Lablab purpureus), Illinois bundleflower (Desman-thus illinoensis), and alfalfa (Medicago sativa) generally resulted in smaller egg population density in soil or number of cysts formed on soybean in the bioassay than the fallow control. Sunn hemp most consistently showed the lowest numbers of eggs and cysts. As a group, legumes resulted in lower egg population densities than monocots, Brassica species, and other dicots.     

Management of Lesion Nematodes and Potato Early Dying with Rotation Crops

Soil-incorporated rotation/green manure crops were evaluated for management of potato early dying caused by Verticillium dahliae and Pratylenchus penetrans. After two years of rotation/green manure and a subsequent potato crop, P. penetrans numbers were less after ‘Saia’ oat/‘Polynema’ marigold, ‘Triple S’ sorghum-sudangrass, or ‘Garry’ oat than ‘Superior’ potato or ‘Humus’ rapeseed. The area under the disease progress curve (AUDPC) for early dying was lowest after Saia oat/marigold, and tuber yields were greater than continuous potato after all crops except sorghum-sudangrass. Saia oat/marigold crops resulted in the greatest tuber yields. After one year of rotation/green manure, a marigold crop increased tuber yields and reduced AUDPC and P. penetrans. In the second potato crop after a single year of rotation, plots previously planted to marigolds had reduced P. penetrans densities and AUDPC and increased tuber yield. Rapeseed supported more P. penetrans than potato, but had greater yields. After two years of rotation/green manure crops and a subsequent potato crop, continuous potato had the highest AUDPC and lowest tuber weight. Rotation with Saia oats (2 yr) and Rudbeckia hirta (1 yr) reduced P. penetrans and increased tuber yields. AUDPC was lowest after R. hirta. Two years of sorghum-sudangrass did not affect P. penetrans, tuber yield or AUDPC. These results demonstrate that P. penetrans may be reduced by one or two years of rotation to non-host or antagonistic plants such as Saia oat, Polynema marigold, or R. hirta and that nematode control may reduce the severity of potato early dying.     

Crop Rotation Effects on Population Densities of Ectoparasitic Nematodes

The influence of rotation crop species on population densities of ectoparasitic nematode species common to soybean and corn fields of the North-Central region of the USA was studied for 5 years in 16 field blocks rotated to corn, soybeans, oats, wheat, and forage mixtures. Each block was sampled each year between mid-July and mid-August. High populations of Helicotylenchus pseudorobustus were maintained in all crops, especially corn. Tylenchorhynchus martini and Xiphinema americanum also developed with all crops, but a second year of corn depressed densities of both species. Paratylenchus projectus densities were high on soybeans and on forage consisting primarily of legumes.     

Managing Nematode Population Densities on Tomato Transplants Using Crop Rotation and a Nematicide

Millet, milo, soybean, crotalaria, and Norman pigeon pea were used in conjunction with clean fallow and a nematicide (fensulfothion) for managing nematode populations in the production of tomato transplants (Lycopersicon esculentum Mill.). Glean fallow was the most effective treatment in suppressing nematode numbers. After 2 years in tomato, root-knot nematodes increased in numbers to damaging levels, and fallow was no longer effective for complete control even in conjunction with fensulfothion. After 4 years in tomato, none of the crops used as summer cover crops alone or in conjunction with fensulfothion reduced numbers of root-knot nematodes in harvested tomato transplants sufficiently to meet Georgia certification regulations. Milo supported large numbers of Macroposthonia ornata and Pratylenchus spp. and crotalaria supported large numbers of Pratylenchus spp. Millet, milo, soybean, crotalaria, and pigeon pea are poor choices for summer cover crops in sites used to produce tomato transplants, because they support large populations of root-knot and other potentially destructive nematodes.     

Management of Heterodera glycines by Cropping and Cultural Practices

Heterodera glycines was identified in North Carolina in 1954, although symptoms of the disease were noted in the state at least 8 years earlier. Crop rotation experiments designed to develop management systems were initiated in 1956. Two or more years in production of a nonhost crop resulted in decreases of the nematode to low or undetectable levels with acceptable subsequent yields of soybean (Glycine max). Because of almost complete dependence on resistant cultivars and (or) nematicides for nematode control, crop rotation experiments were not conducted from 1962 to 1980. Research on control of H. glycines, beginning in 1981, emphasized biological and ecological aspects of the nematode in order to determine cropping systems that restrict the nematode to nondamaging levels. Mortality during embryogenesis was high at temperatures above 30 C. Hatching of eggs occurs readily in May and June. Postinfection development takes 2-3 weeks at weekly mean temperatures of 22-29 C and is slow above and below those temperatures. Egg production is high during the late growing season. Some cultural practices such as planting early maturing cultivars in mid-to-late June and rotation with a nonhost effectively keeps populations at low levels.     

Response of Peanut,Corn, Tobacco,and Soybean to Criconemella ornate

The relative susceptibility of four field crops to Criconemella ornata differed greatly in microplot tests. As few as 178 freshly-introduced C. ornata/500 cm³ of soil stunted peanut. In contrast, this nematode had no effect on the growth of corn or soybean. Large populations remaining after culture of peanut or corn enhanced the growth of tobacco. A problem of comparing the effects of a freshly introduced population of this nematode with large residual populations was encountered. Freshly extracted, greenhouse-grown inoculum caused the typical "yellows disease" on peanut, whereas much greater residual population densities following a poor host (tobacco) had little effect on the growth of peanut. It is suggested that many of the nematodes in the field following a poor host are dead. Peanut supported greater reproduction (up to 970-fold) than did other crops tested. Corn was intermediate, with a population increase as great as 264-fold; soybean and tobacco failed to maintain initial population densities.     

Tropical Rotation Crops Influence Nematode Densities and Vegetable Yields

The effects of eight summer rotation crops on nematode densities and yields of subsequent spring vegetable crops were determined in field studies conducted in north Florida from 1991 to 1993. The crop sequence was as follows: (i) rotation crops during summer 1991; (ii) cover crop of rye (Secale cereale) during winter 1991-92; (iii) ''Lemondrop L'' squash (Cucurbita pepo) during spring 1992; (iv) rotation crops during summer 1992; (v) rye during winter 1992-93; (vi) ''Classic'' eggplant (Solanum melongena) during spring 1993. The eight summer crop rotation treatments were as follows: ''Hale'' castor (Ricinus communis), velvetbean (Mucuna deeringiana), sesame (Sesamum indicum), American jointvetch (Aeschynomene americana), weed fallow, ''SX- 17'' sorghum-sudangrass (Sorghum bicolor x S. sudanense), ''Kirby'' soybean (Glycine max), and ''Clemson Spineless'' okra (Hibiscus esculentus) as a control. Rotations with castor, velvetbean, American jointvetch, and sorghum-sudangrass were most effective in maintaining the lowest population densities of Meloidogyne spp. (a mixture of M. incognita race 1 and M. arenaria race 1), but Paratrichodorus minor built up in the sorghum-sudangrass rotation. Yield of squash was lower (P ≤ 0.05) following sorghum-sudangrass than after any of the other treatments except fallow. Yield of eggplant was greater (P ≤ 0.05) following castor, sesame, or American jointvetch than following okra or fallow. Several of the rotation crops evaluated here may be useful for managing nematodes in the field and for improving yields of subsequent vegetable crops.     

Biology of Thripinema nicklewoodi (Tylenchida), an Obligate Frankliniella occidentalis (Thysanoptera) Parasite

Frankliniella occidentalis, a serious pest of agricultural crops, is difficult to manage because chemical and biological control measures frequently fail to affect F. occidentalis in their preferred microhabitats. Parasitism by the host-specific, entomopathogenic nematode Thripinema nicklewoodi may provide a much-needed alternative to current control strategies. Infection does not cause death of the host; rather, the result is sterilization that leads to suppression of F. occidentalis populations. We describe a simple rearing method and the results from studies aimed at providing details on its biology-both essential first steps to examining its biological control potential. All F. occidentalis life stages are susceptible to infection, but to varying degrees (most susceptible to least susceptible): female pupae, second instar larvae, first instar larvae, male pupae, adult females, adult males. Nematodes emerge from female and male F. occidentalis for approximately 15 and 9 days, with approximately 14 and 7 nematodes emerging per day, respectively. Females and males are short-lived outside of the host, with mean survival rates ranging between 7 and 86 hours. Transmission does not occur in the soil but rather on or within plant structures that are preferred microhabitats visited by F. occidentalis. Results from a dose-response study suggest that augmentative applications of T. nicklewoodi may be useful to generate increased infection rates and subsequent suppression of F. occidentalis populations.     

Review of Pasteuria penetrans: Biology,Ecology, and Biological Control Potential

Pasteuria penetrans is a mycelial, endospore-forming, bacterial parasite that has shown great potential as a biological control agent of root-knot nematodes. Considerable progress has been made during the last 10 years in understanding its biology and importance as an agent capable of effectively suppressing root-knot nematodes in field soil. The objective of this review is to summarize the current knowledge of the biology, ecology, and biological control potential of P. penetrans and other Pasteuria members. Pasteuria spp. are distributed worldwide and have been reported from 323 nematode species belonging to 116 genera of free-living, predatory, plant-parasitic, and entomopathogenic nematodes. Artificial cultivation of P. penetrans has met with limited success; large-scale production of endospores depends on in vivo cultivation. Temperature affects endospore attachment, germination, pathogenesis, and completion of the life cycle in the nematode pseudocoelom. The biological control potential of Pasteuria spp. have been demonstrated on 20 crops; host nematodes include Belonolaimus longicaudatus, Heterodera spp., Meloidogyne spp., and Xiphinema diversicaudatum. Pasteuria penetrans plays an important role in some suppressive soils. The efficacy of the bacterium as a biological control agent has been examined. Approximately 100,000 endospores/g of soil provided immediate control of the peanut root-knot nematode, whereas 1,000 and 5,000 endospores/g of soil each amplified in the host nematode and became suppressive after 3 years.     

Potential of Leguminous Cover Crops in Management of a Mixed Population of Root-knot Nematodes (Meloidogyne spp.)

Root-knot nematode is an important pest in agricultural production worldwide. Crop rotation is the only management strategy in some production systems, especially for resource poor farmers in developing countries. A series of experiments was conducted in the laboratory with several leguminous cover crops to investigate their potential for managing a mixture of root-knot nematodes (Meloidogyne arenaria, M. incognita, M. javanica). The root-knot nematode mixture failed to multiply on Mucuna pruriens and Crotalaria spectabilis but on Dolichos lablab the population increased more than 2- fold when inoculated with 500 and 1,000 nematodes per plant. There was no root-galling on M. pruriens and C. spectabilis but the gall rating was noted on D. lablab. Greater mortality of juvenile root-knot nematodes occurred when exposed to eluants of roots and leaves of leguminous crops than those of tomato; 48.7% of juveniles died after 72 h exposure to root eluant of C. spectabilis. The leaf eluant of D. lablab was toxic to nematodes but the root eluant was not. Thus, different parts of a botanical contain different active ingredients or different concentrations of the same active ingredient. The numbers of root-knot nematode eggs that hatched in root exudates of M. pruriens and C. spectabilis were significantly lower (20% and 26%) than in distilled water, tomato and P. vulgaris root exudates (83%, 72% and 89%) respectively. Tomato lacks nematotoxic compounds found in M. pruriens and C. spectabilis. Three months after inoculating plants with 1,000 root-knot nematode juveniles the populations in pots with M. pruriens, C. spectabilis and C. retusa had been reduced by approximately 79%, 85% and 86% respectively; compared with an increase of 262% nematodes in pots with Phaseolus vulgaris. There was significant reduction of 90% nematodes in fallow pots with no growing plant. The results from this study demonstrate that some leguminous species contain compounds that either kill root-knot nematodes or interfere with hatching and affect their capacity to invade and develop within their roots. M. pruriens, C. spectabilis and C. retusa could be used with effect to decrease a mixed field populations of root-knot nematodes.     

Effects of Pratylenchus penetrans and Rhizoctonia fragariae on Vigor and Yield of Strawberry

Microplot and small field-plot experiments were conducted to determine the effects of Pratylenchus penetrans on strawberry yield over several seasons and to evaluate the effects of nematode control on strawberry vigor and yield. Pratylenchus penetrans alone or in combination with the black root rot pathogen, Rhizoctonia fragariae, reduced strawberry yield in microplots over time. There were no differences in effects on yield among R. fragariae anastomosis groups A, G, or I. The interaction of the two pathogens appeared to be additive rather than synergistic. In field plots infested with P. penetrans alone, plant vigor and yield were increased by the application of carbofuran and fenamiphos nematicides. Nematode control was transitory, as P. penetrans populations were initially suppressed but were not different in samples taken 10 months after treatment. These data highlight the error in associating causality between plant damage and nematode populations based on a correlation of root disease with nematode diagnostic assays from severely diseased plants. These findings may help to explain how nematode numbers can sometimes be higher in healthy plants than in severely diseased plants that lack sufficient roots to maintain nematode populations. Because nematode populations from up to a year before harvest are better correlated with berry yield, preplant nematode diagnostic assays taken a year in advance of harvest may be superior in predicting damage to perennial strawberry yield.     

Pasteuria penetrans for Control of Meloidogyne incognita on Tomato and Cucumber,and M. arenaria on Snapdragon

Meloidogyne incognita and Meloidogyne arenaria are important parasitic nematodes of vegetable and ornamental crops. Microplot and greenhouse experiments were conducted to test commercial formulations of the biocontrol agent Pasteuria penetrans for control of M. incognita on tomato and cucumber and M. arenaria on snapdragon. Three methods of application for P. penetrans were assessed including seed, transplant, and post-plant treatments. Efficacy in controlling galling and reproduction of the two root-knot nematode species was evaluated. Seed treatment application was assessed only for M. incognita on cucumber. Pasteuria treatment rates of a granular transplant formulation ranged from 1.5 × 10⁵ endospores/cm³ to 3 × 10⁵ endospores/cm³ of transplant mix applied at seeding. Additional applications of 1.5 × 10⁵ endospores/cm³ of soil were applied as a liquid formulation to soil post-transplant for both greenhouse and microplot trials. In greenhouse cucumber trials, all Pasteuria treatments were equivalent to steamed soil for reducing M. incognita populations in roots and soil, and reducing nematode reproduction and galling. In cucumber microplot trials there were no differences among treatments for M. incognita populations in roots or soil, eggs/g root, or root condition ratings. Nematode reproduction on cucumber was low with Telone II and with the seed treatment plus post-plant application of Pasteuria, which had the lowest nematode reproduction. However, galling for all Pasteuria treatments was higher than galling with Telone II. Root-knot nematode control with Pasteuria in greenhouse and microplot trials varied on tomato and snapdragon. Positive results were achieved for control of M. incognita with the seed treatment application on cucumber.     

Evaluation of Legumes Common to the Pacific Northwest as Hosts for the Pea Cyst Nematode,Heterodera goettingiana

Seventeen leguminous species common to the Pacific Northwest were evaluated as potential hosts of the pea cyst nematode, Heterodera goettingiana, in both greenhouse and field experiments. In all experiments, juveniles of H. goettingiana penetrated roots of these 17 species with the exception of greenhouse-grown chickpea. Nematodes molted and developed into swollen third-stage or fourth-stage juveniles in many of the plants, but cyst development occurred only in the field on green pea, edible dry pea, and faba bean. More H. goettingiana cysts developed on fava bean than on green pea or edible dry pea. In H. goettingiana-infested soils, cropping sequences that include fava bean and pea should be avoided. However, certain legumes, such as winter vetch, may have the potential of serving as trap crops for H. goettingiana in this region.     

Population Dynamics of Meloidogyne incognita,M. arenaria,and Other Nematodes and Crop Yields in Rotations of Cotton,Peanut, and Wheat Under Minimum Tillage

Wheat, cotton, and peanut were arranged in three cropping sequences to determine the effects of fenamiphos (6.7 kg a.i./ha) and cropping sequence on nematode population densities and crop yields under conservation tillage and irrigation for 6 years. The cropping sequences included a wheat winter cover crop each year and summer crops of cotton every year, peanut every year, or cotton rotated every other year with peanut. The population densities of Meloidogyne spp. and Helicotylenchus dihystera were determined monthly during the experiment. Numbers of M. incognita increased on cotton and decreased on peanut, whereas M. arenaria increased on peanut, and decreased on cotton; both nematode species remained in moderate to high numbers in plots of wheat. Root damage was more severe on cotton than peanut and was not affected by fenamiphos treatment. The H. dihystera population densities were highest in plots with cotton every summer, intermediate in the cotton-peanut rotation, and lowest in plots with peanut every summer. Over all years and cropping sequences, yield increases in fenamiphos treatment over untreated control were 9% for wheat, 8% for cotton, and 0% for peanut. Peanut yields following cotton were generally higher than yields following peanut. These results show that nematode problems may be manageable in cotton and peanut production under conservation tillage and irrigation in the southeastern United States.     

Effects of Switchgrass (Panicum virgatum) Rotations with Peanut (Arachis hypogaea L.) on Nematode Populations and Soil Microflora

A 3-year field rotation study was conducted to assess the potential of switchgrass (Panicum virgatum) to suppress root-knot nematodes (Meloidogyne arenaria), southern blight (Sclerotium rolfsii), and aflatoxigenic fungi (Aspergillus sp.) in peanut (Arachis hypogaea L.) and to assess shifts in microbial populations following crop rotation. Switchgrass did not support populations of root-knot nematodes but supported high populations of nonparasitic nematodes. Peanut with no nematicide applied and following 2 years of switchgrass had the same nematode populations as continuous peanut plus nematicide. Neither previous crop nor nematicide significantly reduced the incidence of pods infected with Aspergillus. However, pod invasion by A. flavus was highest in plots previously planted with peanut and not treated with nematicide. Peanut with nematicide applied at planting following 2 years of switchgrass had significantly less incidence of southern blight than either continuous peanut without nematicide application or peanut without nematicide following 2 years of cotton. Peanut yield did not differ among rotations in either sample year. Effects of crop rotation on the microbial community structure associated with peanut were examined using indices for diversity, richness, and similarity derived from culture-based analyses. Continuous peanut supported a distinctly different rhizosphere bacterial microflora compared to peanut following 1 year of switchgrass, or continuous switchgrass. Richness and diversity indices for continuous peanut rhizosphere and geocarposphere were not consistently different from peanut following switchgrass, but always differed in the specific genera present. These shifts in community structure were associated with changes in parasitic nematode populations.     

An Aphelenchoides sp. nematode Parasitic of Polianthes tuberosa in the Mekong Delta

Polianthes tuberosa is a commercially valuable flower crop in the Mekong Delta of Vietnam that is propagated by the harvesting and planting of bulbs. The cultivation of P. tuberosa is complicated by an endemic nematode infection that damages a high proportion of the plants. Based on morphological criteria and ribosomal RNA gene sequencing, we have determined that the infection is caused by an Aphelenchoides sp. nematode and is most likely Aphelenchoides besseyi. By scoring various parts of harvested plants with flowers for the presence of viable nematodes over a period of six months, we determined that the nematode is an ectoparasite that can survive the intercrop periods, especially in the bulbs that are used to plant new crops. A comparison of farming practices in the Mekong Delta failed to identify useful control methods, but rather indicated that fields that have cultivated P. tuberosa for the longest periods suffer the worst damage from the nematode infection. Finally, we demonstrated that the nematode is capable of infecting 30 rice cultivars but does not cause the white tip disease usually associated with A. besseyi infection.     

Problems and Strategies Associated with Long-term Use of Nematode Resistant Cultivars

Plant-parasitic nematodes are obligate parasites, and planting cultivars that are highly resistant to these organisms places extensive selection pressure on the target species and affects nontarget nematodes as well. Problems encountered with long-term planting of cultivars resistant to nematodes include shifts in nematode races or species and the occurrence of multiple species of nematodes within the same field. These problems can be alleviated to some extent when crop management is used to lessen the selection pressure for change on the nematode populations. Race shifts within populations and possibly shifts between nematode species can be delayed by rotating susceptible cultivars and nonhost crops with resistant cultivars. Nematicides in conjunction with resistant cultivars may be used to limit damage by multiple species of nematodes. Some cultivars have resistance to multiple species of nematodes, but greatly increased research effort is needed in this area. More intensive plant breeding effort will be required to make nematode resistant cultivars competitive in quality and yield with more productive, susceptible cultivars.